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in 1957, JBS Haldane calculated, under a certain set of assumptions, the rate at which a new beneficial mutation can reach fixations in a population. That rate was about 1 every 300 generations. Haldane felt that this was problematic, for reasons I will not get into here.
I bring this up because this was unfortunately referred to as "Haldane's dilemma", and was pounced upon in the 1990s by creationist electrical engineer Walter ReMine in a book that he wrote. He used Haldane's model to conclude that at most, 1667 beneficial mutations could have been fixed in the lineage leading to humans since the split with the lineage leading to chimps. He extrapolated that 1667 fixed beneficial mutations was just not enough to account for human evolution from a common ancestor with chimps (despite never even suggesting which new traits were needed that the common ancestor did not have). He never actually explained why this was a problem, except with the standard 'argument via awe and big numbers' (which consisted entirely of rhetorical questions, like even if there 500,000 such mutations, would even that be enough to get a "sapien from a simian?" Yes, that was supposed to be an argument).
Well, this "dilemma" came up recently in a thread discussing a paper from creationist John Sanford et al's relatively recent paper in which these creationists declare that there is just not enough time for evolution to have occurred (providing that evolution occurs via a series of pre-specified mutations occurring one at a time and one after another in regulatory sequence...), and I was reminded of the habit creationists have of providing quotes from people only as it suits them.
ReMine, for example, relied heavily on geneticist Warren Ewens as his primary resource for his pro-Haldane dilemma argument. He provided many quotes from Ewens in which it seemed that he agreed that Haldane's model was indeed a 'problem' for evolution as it limited the number of beneficial mutations that could become fixed over time.
And yes, Ewens did believe that Haldane's model, as calculated by Haldane, was a problem.
But more importantly, Ewens did not think Haldane's model was accurate, and thus there is no "dilemma" - but Remine just, I guess, forgot to mention that.
Here is Ewens, in an interview in 2004:
And further:
AP: Can I follow that up? Can you, in layman’s terms, explain why you think that there is no upper limit in the way that Haldane suggested?
WE: I can, but it becomes rather mathematical. Let me approach it this way. Suppose that you consider one gene locus only, at which a superior allele is replacing an inferior allele through natural selection. In broad terms, what this requires is that individuals carrying the superior allele have on average somewhat more offspring than the mean number of offspring per parent, otherwise the frequency of the superior allele would not increase. This introduces a concept of a “one-locus substitutional load,” and a formal numerical value for this load is fairly easily calculated. However, the crux of the problem arises when one considers the many, perhaps hundreds or even thousands, substitution processes that are being carried out at any one time. In his mathematical treatment of this “multi-locus” situation, Kimura, for example, in effect simply multiplied the loads at the various individual substituting loci to arrive at an overall total load. The load so calculated was enormous. This uses a reductionist approach to the load question, and to me, this reductionist approach is not the right way of doing things. Further, the multiplicative assumption is, to me, unjustified. It is the selectively favored individuals, carrying a variety of different genes at different loci, who are reproducing and being required to contribute more offspring than the average. If you consider load arguments from that individual-based, non-reductionist basis, the mathematical edifice which Kimura built up just evaporates, and in my view the very severe load calculations which he obtained by his approach became irrelevant and misleading. The individual-based calculations that I made indicated to me that there is no unbearable substitutional load.
ReMine's book came out in 1993. I find it hard to believe that Ewens did not already have these same opinions and had not already published the relevant papers well before then (most of his relevant papers on this issue came out in the 1970s). Not hard to believe that he told this all to ReMine and ReMine, being a creationist with souls to save and profits from a book to earn*, just ignored it all because it did not help his cause.
*Another of ReMine's (and other creationists') tactics is to artificially inflate the relevance and impact of their books to not only boost their sales, but to boost their credibility (for the cause, after all). ReMine did this by sending copies, unsolicited, to public and college libraries, or sending boxes of the book to fans and having them 'donate' the book to the same. Thus, he could then boast that his book is on the shelves of major universities at the like. I have seen 2 copies of ReMine's book at 2 of the universities that I have been associated with - both were "gifts" (i.e., the university did not actually purchase them).
I bring this up because this was unfortunately referred to as "Haldane's dilemma", and was pounced upon in the 1990s by creationist electrical engineer Walter ReMine in a book that he wrote. He used Haldane's model to conclude that at most, 1667 beneficial mutations could have been fixed in the lineage leading to humans since the split with the lineage leading to chimps. He extrapolated that 1667 fixed beneficial mutations was just not enough to account for human evolution from a common ancestor with chimps (despite never even suggesting which new traits were needed that the common ancestor did not have). He never actually explained why this was a problem, except with the standard 'argument via awe and big numbers' (which consisted entirely of rhetorical questions, like even if there 500,000 such mutations, would even that be enough to get a "sapien from a simian?" Yes, that was supposed to be an argument).
Well, this "dilemma" came up recently in a thread discussing a paper from creationist John Sanford et al's relatively recent paper in which these creationists declare that there is just not enough time for evolution to have occurred (providing that evolution occurs via a series of pre-specified mutations occurring one at a time and one after another in regulatory sequence...), and I was reminded of the habit creationists have of providing quotes from people only as it suits them.
ReMine, for example, relied heavily on geneticist Warren Ewens as his primary resource for his pro-Haldane dilemma argument. He provided many quotes from Ewens in which it seemed that he agreed that Haldane's model was indeed a 'problem' for evolution as it limited the number of beneficial mutations that could become fixed over time.
And yes, Ewens did believe that Haldane's model, as calculated by Haldane, was a problem.
But more importantly, Ewens did not think Haldane's model was accurate, and thus there is no "dilemma" - but Remine just, I guess, forgot to mention that.
Here is Ewens, in an interview in 2004:
A second form of the load concept was introduced by the British biologist-mathematician Haldane who claimed, in 1957, that substitutions in a Darwinian evolutionary process could not proceed at more than a certain comparatively slow rate, because if they were to proceed at a faster rate, there would be an excessive “substitutional load.” Since Haldane was so famous, that concept attracted a lot of attention. In particular, Crow and Kimura made various substitutional load calculations around 1960, that is at about that time that I was becoming interested in genetics.
Perhaps the only disagreement I ever had with Crow concerned the substitutional load, because I never thought that the calculations concerning this load, which he and others carried out, were appropriate. From the very start, my own calculations suggested to me that Haldane’s arguments were misguided and indeed erroneous, and that there is no practical upper limit to the rate at which substitutions can occur under Darwinian natural selection.
Perhaps the only disagreement I ever had with Crow concerned the substitutional load, because I never thought that the calculations concerning this load, which he and others carried out, were appropriate. From the very start, my own calculations suggested to me that Haldane’s arguments were misguided and indeed erroneous, and that there is no practical upper limit to the rate at which substitutions can occur under Darwinian natural selection.
And further:
AP: Can I follow that up? Can you, in layman’s terms, explain why you think that there is no upper limit in the way that Haldane suggested?
WE: I can, but it becomes rather mathematical. Let me approach it this way. Suppose that you consider one gene locus only, at which a superior allele is replacing an inferior allele through natural selection. In broad terms, what this requires is that individuals carrying the superior allele have on average somewhat more offspring than the mean number of offspring per parent, otherwise the frequency of the superior allele would not increase. This introduces a concept of a “one-locus substitutional load,” and a formal numerical value for this load is fairly easily calculated. However, the crux of the problem arises when one considers the many, perhaps hundreds or even thousands, substitution processes that are being carried out at any one time. In his mathematical treatment of this “multi-locus” situation, Kimura, for example, in effect simply multiplied the loads at the various individual substituting loci to arrive at an overall total load. The load so calculated was enormous. This uses a reductionist approach to the load question, and to me, this reductionist approach is not the right way of doing things. Further, the multiplicative assumption is, to me, unjustified. It is the selectively favored individuals, carrying a variety of different genes at different loci, who are reproducing and being required to contribute more offspring than the average. If you consider load arguments from that individual-based, non-reductionist basis, the mathematical edifice which Kimura built up just evaporates, and in my view the very severe load calculations which he obtained by his approach became irrelevant and misleading. The individual-based calculations that I made indicated to me that there is no unbearable substitutional load.
ReMine's book came out in 1993. I find it hard to believe that Ewens did not already have these same opinions and had not already published the relevant papers well before then (most of his relevant papers on this issue came out in the 1970s). Not hard to believe that he told this all to ReMine and ReMine, being a creationist with souls to save and profits from a book to earn*, just ignored it all because it did not help his cause.
*Another of ReMine's (and other creationists') tactics is to artificially inflate the relevance and impact of their books to not only boost their sales, but to boost their credibility (for the cause, after all). ReMine did this by sending copies, unsolicited, to public and college libraries, or sending boxes of the book to fans and having them 'donate' the book to the same. Thus, he could then boast that his book is on the shelves of major universities at the like. I have seen 2 copies of ReMine's book at 2 of the universities that I have been associated with - both were "gifts" (i.e., the university did not actually purchase them).
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